Foraging mode and evolution of strike-induced chemosensory searching in lizards

Strike-induced chemosensory searching (SICS) in lizards and snakes is a means of relocating prey by scent-trailing. The two main components of SICS are an elevated tongue-flick rate for vomerolfactory sampling after biting prey (PETF) and searching movements. In combination, these behaviors permit scent-trailing. Prey chemical discrimination, which is a prerequisite for SICS, is present in active foragers, but not in ambush foragers. Using comparative data, I show that searching movements and SICS have undergone correlated evolution with foraging mode and with prey chemical discrimination in lizards. This suggests that active foraging selects for prey chemical discrimination, which is then employed to search for escaped prey using the typical movements and tongue-flicking behaviors of active foragers. SICS in lizards is simply heightened active foraging after biting prey. In nonvenomous snakes, SICS is similar to that in lizards but is not restricted to active foragers. Only highly venomous snakes voluntarily release dangerous prey upon envenomation, pause to let the venom incapacitate the prey, and then relocate the prey by scent-trailing. PETF was observed in two ambush foragers and is not evolutionarily correlated with foraging mode or searching movements. Because it occurs in species lacking prey chemical discrimination, such PETF may be a response to gustatory cues or to internal chemicals not encountered on surfaces or trails of uninjured prey.     

Elevation in tongue-flick rate after biting prey in the broad-headed skink,Eumeces laticeps

A postbite elevation in tongue-flicking (PETF) rate occurs in adult male broad-headed skinks,Eumeces laticeps. Males having bitten neonatal mice showed significantly higher tongue-flicking rates in the 2 min following experimental removal of the prey than did males in several control conditions. In a second experiment designed to separate the effects of tactile and chemical stimulation of the oral cavity during biting, males tongue-flicked at significantly higher rates in response to swabs bearing surface chemicals from neonatal mice than to identical swabs lacking the surface chemicals. These findings agree with previous data showing that PETF and searching movements occur in those families of lizards that can detect prey chemicals and use the tongue to do so during active foraging. The occurrence of PETF and putative searching movements supports the interpretation that PETF represents an attempt to relocate lost prey by chemosensory means. PETF was much briefer inE. laticeps than in many snakes and in a representative species from another lizard family and was not detectable after the first minute. This brevity is consistent with the prediction that PETF should be brief in squamates that feed on prey not likely to be located by scent-trailing.     

Prolonged poststrike elevation in tongue-flicking rate with rapid onset in gila monster,Heloderma suspectum: Relation to diet and foraging and implications for evolution of chemosensory searching

Experimental tests showed that poststrike elevation in tongue-flicking rate (PETF) and strike-induced chemosensory searching (SICS) in the gila monster last longer than reported for any other lizard. Based on analysis of numbers of tongue-flicks emitted in 5-min intervals, significant PETF was detected in all intervals up to and including minutes 41–45. Using 10-min intervals, PETF lasted though minutes 46–55. Two of eight individuals continued tongue-flicking throughout the 60 min after biting prey, whereas all individuals ceased tongue-flicking in a control condition after minute 35. The apparent presence of PETF lasting at least an hour in some individuals suggests that there may be important individual differences in duration of PETF. PETF and/or SICS are present in all families of autarchoglossan lizards studied except Cordylidae, the only family lacking linguallly mediated prey chemical discrimination. However, its duration is known to be greater than 2-min only in Helodermatidae and Varanidae, the living representatives of Varanoidea. That prolonged PETF and SICS are typical of snakes provides another character supporting a possible a varanoid ancestry for Serpentes. Analysis of 1-min intervals showed that PETF occurred in the first minute. A review of the literature suggests that a pause in tongue-flicking and delay of searching movements are absent in lizards and the few nonvenomous colubrid snakes tested. The delayed onset of SICS may be a specific adaptation of some viperid snakes to allow potentially dangerous prey to be rendered harmless by venom following voluntary release after envenomation and preceding further physical contact with the prey.     

Discriminative Response to Animal, But Not Plant, Chemicals by an Insectivorous, Actively Foraging Lizard, Scincella lateralis, and Differential Response to Surface and Internal Prey Cues

Responses by the insectivorous, actively foraging scincid lizard, Scincella lateralis, to chemical cues from a plant food favored by herbivorous lizards, its ability to discriminate prey chemicals from control substances, and its relative response to internal and surface prey chemicals were studied experimentally. We presented chemical cues to the lizards on cotton swabs and recorded their tongue-flicks and biting attacks on the swabs. The lizards exhibited significantly greater tongue-flick rates and biting frequencies to prey surface cues than to plant surface chemicals from romaine lettuce, diluted cologne (pungency control), and deionized water. Responses to the plant stimuli did not differ from those to the two control stimuli, in contrast with strong responses to the same plant cues by herbivores. This finding provides the first information suggesting that chemosensory response may be adapted to diet, with responsiveness to plant stimuli evolving de novo in herbivores. Biting and tongue-flicking responses were significantly greater to cricket chemicals than to all other stimuli, among which there were no differences. Thus, the lizards are capable of prey chemical discrimination, which may be ubiquitous among actively foraging lizards. The lizards exhibited more frequent biting and higher tongue-flick rates to internal than surface prey chemicals. Although different methods of stimulus preparation are appropriate for different purposes, we conclude that prey surface chemicals available to foraging lizards are most desirable for studies bearing on location and identification of prey.     

Postbite elevation in tongue-flicking rate by an iguanian lizard,Dipsosaurus dorsalis

The herbivorous iguanid lizardDipsosaurus dorsalis exhibited PETF (postbite elevation in tongue-flicking rate), an increase in tongue-flicking rate after experimental removal from the mouth of food that had been bitten. This was demonstrated by a significantly higher tongue-flick rate after having bitten food than in three experimental conditions controlling for responses to the experimental setting, sight of food, and mechanical disturbance caused by the experimental removal of food from a lizard's mouth. As in most other families of lizards, PETF was brief, occurring only during the first minute. Lizards are divided into two major suprafamilial taxa, Iguania and Scleroglossa, consisting of carnivorous species characterized by two major foraging modes, ambush and active, and of herbivores and omnivores. PETF is absent in the two families of carnivorous iguanian lizards studied that are ambush foragers but present in three families of scleroglossan lizards that are active foragers. However, PETF is absent in the two species studied in a scleroglossan family, Gekkonidae, which forages by ambush, and present in an iguanian herbivore, as reported herein. We propose that the presence or absence of PETF, in addition to its phylogenetic determinants, is adaptively adjusted to foraging mode.     

Chemical discrimination by tongue-flicking in lizards: A review with hypotheses on its origin and its ecological and phylogenetic relationships

Tongue-flicking is a synapomorphy of squamate reptiles functioning to sample chemicals for vomerolfactory analysis, which became possible in primitive squamates when ducts opened from the vomeronasal organs to the roof of the mouth. Extant iguanian lizards in families that do not use the tongue to sample chemical prey cues prior to attack partially protrude it in two feeding contexts: during capture by lingual prehension and after oral contact with prey. These lizards do not exhibit strike-induced chemosensory searching. Lingual prey prehension is present in iguanian lizards and inSphenodon, the sister taxon of Squamata. During attempts to capture prey, the tongues of primitive squamates inevitably made incidental contact with environmental substrates bearing chemicals deposited by prey, conspecifics, and predators. Such contact presumably induced selection for tongue-flicking and ability to identify biologically important chemicals. Most iguanian lizards are ambush foragers that use immobility as a major antipredatory defense. Because tongue-flicking at an ambush post would not allow chemical search beyond the vicinity of the head and would render them easier for predators and prey to detect, typical iguanians tongue-flick neither while foraging nor to identify predators. They do detect pheromones by tongue-flicking. Scleroglossan lizards are typically active foragers that rely on speed to escape. Being freer to move the tongue, they have evolved lingual sampling allowing detection of chemical cues of conspecifics, predators, and prey, as well as strike-induced chemosensory searching, some can follow pheromone trails by tongue-flicking. Some families have lingual morphology and behavior specialized for chemosensory sampling. In varanids and snakes, the taxa showing the greatest lingual specialization, additional prey-related chemosensory behaviors have evolved. In iguanian and scleroglossan families that have secondarily adopted the foraging mode typical of the other taxon, prey chemical discrimination involving tongue-flicking and strike-induced chemosensory searching are typical for the foraging mode rather than the taxon. Because foraging mode and state of prey chemical discrimination are stable within squamate families and to a large extent in higher taxa, both features have been retained from the ancestral condition in most families. However, in three cases in which foraging mode has changed from its ancestral state, the state of prey chemical discrimination has also changed, indicating that prey chemical discrimination is adaptively adjusted to foraging mode. Indeed, acquisition of lingually mediated prey chemical discrimination may have made feasible the evolution of active foraging, which in turn appears to have profoundly influenced the further evolution of squamate chemosensory structures and behavior, placing a selective premium on features enhancing the tongue's efficiency as a chemical sampling device. The advent of tongue-flicking to sample prey chemicals and thus detect hidden prey may have allowed generalist (cruise) or ambush foragers, if early squamates were such, to become specialists in active foraging. Alternatively, if the common ancestors of squamates were active foragers, the adoption of ambush foraging would have selected against participation of the tongue in locating prey. Acting jointly, tongue-flicking and active foraging have had momentous consequences for squamate diversification. Specialization for active foraging would appear to have had ramifying effects on antipredatory defenses, body form, territoriality, mating systems, and reproductive physiology.     

Correspondence Between Diet and Food Chemical Discriminations by Omnivorous Geckos (Rhacodactylus)

Chemosensory responses to food are correlated with geographic variation in diet of some colubrid snakes, but the influence of diet on chemosensory behavior has not been established generally in snakes or lizards. Most lizards are generalist predators of small animals, making it difficult to study effects of diet, but herbivory and omnivory have evolved in several lineages, providing an excellent opportunity to study the effects of dietary change on chemosensory behavior. Based on ecological considerations, I argue that inclusion of plants in the diet of lizards that evolved from ambush foragers lacking prey chemical discrimination might be expected to evolve responsiveness to plant food chemicals. If animal prey also are retained in the diet, then responsiveness to prey chemicals should evolve as well. I experimentally studied tongue-flicking and biting responses by omnivorous geckos of the genus Rhacodactylus to chemical stimuli from plant and animal foods and control substances presented on cotton swabs. The lizards exhibited significantly greater responses to plant stimuli than to control stimuli. One of two species tested responded strongly to cricket chemicals, but the other showed no significant response to mouse surface chemical stimuli. The results support the hypothesis that dietary shifts induce corresponding changes in chemosensory response, but establishment of correlated evolution between diet and food chemical discriminations in lizards will require study of many herbivores/omnivores and insectivores as controls.     

Chemical discrimination of prey by naive neonate Gould's monitorsVaranus gouldii

Previous studies have demonstrated that actively foraging autarchoglossan lizards rely in part on chemoreception to detect and locate prey. In one of two experiments, neonate Gould's monitorsVaranus gouldii were studied to determine whether they were able to discriminate between multiple prey odors and control odors by tongue-flicking. Responses of lizards to deionized water, a pungency control (cologne), mouse, gecko, and cricket odors on cotton-tipped applicators were studied in experiments using repeated-measures designs and using the tongue-flick attack score (TFAS) as the primary measure of response strength. The TFAS was greater in response to cricket odors than to other prey odors or to either of the control stimuli, and there was no statistically significant difference in response between control stimuli. Range of tongue-flicks elicited by cricket odor were greater than those for other prey odors and control stimuli. Only applicators bearing cricket odor were bitten. In the second experiment, lizards were tested to determine whether they respond differently to chemical stimuli taken from the exoskeleton vs. internal fluids of crickets. TFAS were slightly higher for chemical stimuli taken from internal fluids, but not significantly so. Lizards bit applicators in both conditions. Details of responses to experimental trials are discussed in relation to the feeding behavior of this species.     

Strike-induced chemosensory searching occurs in lizards

Strike-induced chemosensory searching (SICS), previously known only in snakes, is experimentally demonstrated in a lizard,Varanus exanthematicus. Tongue-flicking rate was significantly greater after striking the prey than following three control conditions. The occurrence of SICS in a varanid lizard suggests that SICS may serve to help relocate dropped or escaped prey not only in snakes, but in other squamates that use the tongue as a chemosensory sampling device during foraging. This in turn suggests the need for further studies of the taxonomic distribution of SICS in squamates and of its relationship to tongue use during foraging and feeding.     

Discrimination of Prey, But Not Plant, Chemicals by Actively Foraging, Insectivorous Lizards, the Lacertid Takydromus sexlineatus and the Teiid Cnemidophorus gularis

Sampling environmental chemicals to reveal prey and predators and to provide information about conspecifics is highly developed in lizards. Actively foraging lizards can discriminate between prey chemicals and control stimuli, but ambush foragers do not exhibit prey chemical discrimination. Recent experiments on a few species of herbivorous lizards have also demonstrated an ability to identify plant food chemicals. We studied chemosensory responses to chemicals from prey and palatable plants in two species of actively foraging, insectivorous lizards. Both the lacertid Takydromus sexlineatus and the teiid Cnemidophorus gularis exhibited strong responses to prey chemicals, but not to plant chemicals. These findings increase confidence in the relationship between prey chemical discrimination and foraging mode, which is based on data for very few species per family. They also provide data showing that actively foraging insectivores in two families do not respond strongly to plant cues. Such information is essential for eventual comparative studies of the relationship between plant diet and responses to food chemicals. The traditional method of presenting stimuli by using hand-held cotton swabs worked well for T. sexlineatus but could not be used for C. gularis due to repeated escape attempts. When stimuli were presented to C. gularis on ceramic tiles and no experimenter was visible, the lizards responded readily. Presentation of stimuli on tiles in the absence of a visible experimenter may be a valuable approach to study of food chemical discrimination by active foragers in which antipredatory behavior interferes with responses to swabs.     

Naive ophiophagus lizards recognize and avoid venomous snakes using chemical cues

Monitor lizards prey on snakes. Conversely, venomous snakes prey on juvenile monitor lizards. Immediately after hatching, monitor lizards are naive to all prey items, thus correct assessment of snake prey is paramount for survival. Experiments were conducted to determine how hatchling monitor lizards (Varanus albigularis) with no previous exposure to snakes reacted to sympatric venomous and nonvenomous snakes. Hatchling lizards attacked harmless snakes, but avoided venomous species. Lizards readily accepted meat from skinned snakes, regardless of species. When invertebrate prey covered with skin segments from venomous snakes were restrained from moving, they were usually investigated by tongue-flicking and rejected. Unrestrained skin-covered prey, however, were generally attacked and eaten without prior evaluation by tongue-flicking. Attack was inhibited in trials in which unrestrained prey were tongue-flicked, suggesting that chemical cues contained in snake skins mediate avoidance of venomous snakes. Selection for the ability to perceive snake integumental chemicals may be especially strong in species that both consume and are consumed by snakes.     

Responses to major categories of food chemicals by the lizard Podarcis lilfordi

Many lizards are capable of identifying food using only chemical cues from food surfaces, but almost nothing is known about the types of compounds that are effective stimuli. We experimentally studied lingual and biting responses by a lacertid lizard, Podarcis lilfordi, to single representatives of three major categories of food chemicals, sucrose as a carbohydrate, pure pork fat as a mixture of lipids, and bovine gamma globulin as a protein. In 60-sec trials in which stimuli were presented on cotton swabs, the lizards detected all three stimuli, exhibiting more tongue-flicks, licks, or bites, or a greater tongue-flick attack score (TFAS; overall measure of response strength to prey stimuli) than to deionized water. The initial response to all stimuli was tongue-flicking, but the lizards discriminated among the types of chemical stimuli. After preliminary tongue-flicks, the lizards responded to sucrose solutions by licking at high rates, to pure pork fat by biting, and to protein by a combination of additional tongue-flicks and biting. Biting is a feeding response to prey or solid plant material. Licking is a feeding response to sugars in nectar or ripe fruit. Its frequency increased with sucrose concentration. Our data suggest that lizards can identify several types of chemicals associated with food and direct feeding attempts to sources of such chemicals in the absence of visual cues.     

Response of brown tree snakes (Boiga irregularis) to human blood

Ten specimens ofBoiga irregularis were presented with clean or bloody tampons. The latter were used by women during menses. Trial duration was 60 sec, intertrial interval was 24 hr, and the dependent variable was rate of tongue flicking (a measure of chemosensory investigation). Bloody tampons elicited significantly more tongue flicking than did control tampons. An additional snake is shown attacking and ingesting a soiled tampon, confirming that chemosensory interest was associated with predatory behavior.     

Prey detection by vomeronasal chemoreception in a plethodontid salamander

While chemoreception is involved in a wide variety of salamander behaviors, the chemosensory system that mediates specific behaviors is rarely known. We investigated the role of the vomeronasal system (VNS) in foraging behavior of the red-backed salamander (Plethodon cinereus) by manipulating salamanders' abilities to detect nonvolatile chemical cues emitted by potential prey. Subjects received one of three treatments: (1) impaired vomeronasal system, (2) sham manipulation, and (3) no manipulation. The role of the VNS in mediating foraging on motile prey (Drosophila melanogaster) was investigated under three light conditions (bright, dim, dark). Salamanders with impaired VNSs foraged less efficiently than either of the other experimental groups by displaying the longest latency to attack and the lowest rate of prey capture, especially in the absence of visual cues. A second experiment utilized freshly killed prey to determine whether the VNS takes on added importance in the absence of visual or tactile cues associated with moving prey. Animals with impaired VNSs showed a decreased foraging efficiency on stationary prey under both dark and light conditions. In addition, a mark–recapture study of VNS-impaired and sham salamanders in the field also indicated that salamanders with impaired VNSs consumed fewer stationary prey compared to shams. The study indicates that the VNS plays a substantial role in the foraging behavior of the plethodontid salamander, P. cinereus.     

Tongue-flicking and biting in response to chemical food stimuli by an iguanid lizard (Dipsosaurus dorsalis) having sealed vomeronasal ducts: Vomerolfaction may mediate these behavioral responses

In the iguanid lizardDipsosaurus dorsalis, chemical food stimuli were discriminated from other odorants by vomerolfaction. This was demonstrated in a 2 × 3 experiment in which groups of lizards with sealed vomeronasal ducts or sham-sealed vomeronasal ducts responded to carrot chemical stimuli, cologne, and distilled water presented on cotton-tipped applicators. Abilities to detect and discriminate food chemicals were abolished in lizards having sealed vomeronasal ducts. For tongue-flick attack score and number of lizards biting, the sham-sealed group responded more strongly to carrot stimuli than to the control stimuli, but the group having sealed ducts did not. Lizards having sham-sealed ducts responded more strongly to carrot stimuli than did lizards having sealed ducts; responses by the two groups of lizards to control stimuli did not differ. Tongue-flicking occurred when the vomeronasal system detected a chemical stimulus from either carrot or cologne. Biting occurred only when the vomeronasal organ detected food stimuli (from carrot). Most duct-sealed lizards opened their mouths, some repeatedly. Mouth-opening thus occurs when the vomeronasal organ does not detect chemicals. It may be an attempt to stimulate or prime the vomeronasal organ or to dislodge the sealant.     

Evaluation of Swab and Related Tests as a Bioassay for Assessing Responses by Squamate Reptiles to Chemical Stimuli

The ability of squamates to detect chemical cues from adaptively important sources including prey, predators, and conspecifics has been tested frequently by presenting stimuli on cotton-tipped swabs or ceramic tiles. In many such studies the primary response variable is tongue-flicking, which is widely interpreted to indicate sampling for vomerolfaction. I review the basic experimental method and consider limitations regarding its application and interpretation and ways to overcome them. Effects of experimenter proximity and the assumed invisibility of chemical stimuli are considered, as are use of cologne as a pungency control, senses used in making chemical discriminations, and interpretation of results when there are no significant response differences among stimulus classes. Although the assumption that tongue-flicking reveals vomerolfactory sampling and the necessity of an intact vomeronasal system for normal responses to pheromones have been demonstrated where tested, very few species have been examined. In some squamates for which these assumptions have not been examined experimentally, especially eublepharid geckos, attacks on swabs bearing prey chemicals and performance of antipredatory displays in response to predator chemicals occur with no prior tongue-flicking. Not only are assays based on tongue-flicking useless in such cases, but the discriminations are likely based on olfaction. Issues specific to the study of responses to prey chemicals, predator chemicals, and pheromones are discussed. For many purposes, swab tests provide rapid, conclusive assays of ability to respond differentially to biologically relevant stimuli. However, other methods may be superior for studying some responses, and swab tests are not always applicable.     

Experimental Evidence of an Age-Specific Shift in Chemical Detection of Predators in a Lizard

The risk posed by predation is one of the most fundamental aspects of an animal's environment. Avoidance of predators implies an ability to obtain reliable information about the risk of predation, and for many species, chemosensory cues are likely to be an important source of such information. Chemosensory cues reliably reveal the presence of predators or their presence in the recent past. We used retreat site selection experiments to test whether the Australian scincid lizard Eulamprus heatwoleiM uses chemical cues for predator detection and avoidance. Both adult and juvenile lizards were given the choice of retreat sites treated with scents from invertebrate predators, as well as sympatric and allopatric snake predators. Some of the snake predators were known to eat E. heatwolei, while others did not pose a predation threat. All invertebrate predators posed a risk to juveniles, but not adults because of their size. We found that juvenile E. heatwolei avoided predator odors more strongly than adults. Juveniles avoided both invertebrate predators and snakes, and the strongest response was toward the funnelweb spider, the only ambush predator used in this experiment. This result may demonstrate the importance of predator ecology in the evolution of predator detection mechanisms, with chemical cues being more useful in detecting sedentary predators than active predators. Adult lizards showed no avoidance behavior toward predator odors. This result suggests an age specific shift in predator avoidance behavior as lizards get older and become too large for many predators. However, adults showed no response to the odor from the red-bellied black snake, a known predator of adult E. heatwolei. This finding further demonstrates the importance of predator ecology when examining communication between predators and prey. Chemical cues, which are persistent long after predators have vacated the area, may not be useful in detecting the red-bellied black snake, a wide-ranging active forager.     

Chemicals of predatory mosquitofish (Gambusia affinis) influence selection of oviposition site by Culex mosquitoes

Ovipositing insects may avoid aquatic sites where there is high predation risk to their offspring, but the proximate mechanisms that mediate avoidance behavior are poorly resolved. We conducted an experiment to determine whether mosquitoes would reduce oviposition rates in pools containing chemicals of the mosquitofish (Gambusia affinis), a voracious predator that is widely employed to control mosquitoes. Experimental treatments consisted of outdoor pools that contained known concentrations of fish chemicals (low, medium, or high) or no fish chemicals (control). The pools were arranged in a randomized block design, and the number of mosquito larvae in each pool served as the response variable to estimate relative oviposition rate. Members of the Culex pipiens complex were the main colonizers of the pools. The mean number of larvae per pool differed among treatments (P = 0.026) and was about three times greater in control pools compared with those receiving medium and high concentrations of fish chemicals. Pairwise comparisons indicate that only medium and high treatments differed significantly from controls, suggesting that a threshold concentration exists below which mosquitoes cannot reliably detect predators. Our data suggest that the effectiveness of Gambusia affinis in controlling mosquitoes may be compromised if adult mosquitoes respond to fish stocking by shifting to nearby breeding sites that lack fish. We discuss issues concerning the use of Gambusia in biological control programs within the context of these new findings.     

Feeding stimulatory and inhibitory chemicals from an acceptable nonhost plant forManduca sexta: Improved detection by larvae deprived of selected chemosensory organs

The chemical basis of feeding responses to the acceptable nonhost plantVigna sinensis (cowpea) by larvae ofManduca sexta was investigated using chemical isolation techniques directed by a novel chemosensory-based bioassay. The presence of feeding stimulatory and inhibitory compounds in leaves or leaf extracts was determined in a two-choice preference test using leaf disks or glass fiber filter paper disks laced with leaf extract as test substrate and filter paper disks laced with water as control. Larvae strongly prefer the control disks over leaf disks, indicating the presence of feeding inhibitory compounds in the leaf. An ethanol extract of both fresh and dried leaves neither stimulated nor inhibited feeding. The cause of this inactivity was examined by using larvae that respond strongly to either feeding stimulatory or inhibitory compounds due to selective chemosensory deprivation. Larvae having chemosensory organs remaining only on the maxillary palps are stimulated to feed by whole leaf disks and by the ethanol extracts. In contrast, larvae having only the medial and lateral maxillary sensilla styloconica and the epipharyngeal sensilla remaining are strongly inhibited by whole leaf disks and the ethanol extract of fresh leaves. Thus, the ethanol extract contains both feeding stimulatory and inhibitory compounds, which elicit opposite behavioral effects in unoperated larvae, therefore nullifying any stimulatory and inhibitory activity. These compounds can only be demonstrated by using discrimination-enhanced larvae in the choice tests. Further isolation of the feeding stimulatory principle inV. sinensis yielded two separate fractions of neutral compounds, suggesting at least two different chemicals belonging to two different classes: nonpolar and polar lipids. Feeding inhibitory chemicals have apparently polar properties because strong activity was found in the ethyl acetate and methanol extracts of dried leaves. The role of feeding stimulatory and inhibitory compounds in food selection ofM. sexta larvae is discussed.     

Predatory behavior inTupinambis teguixin (Sauria: Teiidae). I. Tongue-flicking responses to chemical food stimuli

Black tegu lizards (Tupinambis teguixin) have the ability to detect food odors and discriminate between them and nonfood odors. This was tested by offering chemical stimuli on cotton-tipped applicators to the animals. Stimuli were from two plant and two animal species known to be principal items in these lizards' diets, demineralized water as an odorless control, and eaude-cologne as an odorous control lacking feeding or social importance. Tongueflick attack score, latency to attack, preattack tongue-flicks, and number of attacks were analyzed. The results clearly demonstrated that this species responds to chemical food stimuli, but does not respond to odorless nonfood stimuli. Responses differed among food types. There were no sex differences. These results are in agreement with the prediction that lizards having forked tongues and an active foraging mode rely on chemical cues for feeding.